Harrison’s Rule Corroborated Among Nomadine Cuckoo Bees (Hymenoptera: Apidae: Nomadinae) : The Role of Body Size in Host-Switching Dynamics

Abstract

Harrison’s rule, that body size is positively correlated between parasites and hosts, has been reported in a range of taxa, but whether the rule is applicable to cleptoparasitic insects is poorly understood. Subfamily Nomadinae, the largest group of cleptoparasitic bees, usurp the nests of a variety of host bees. Within the subfamily, Nomada exploits the most diverse hosts, using at least ten genera from five families. Not only they use diverse hosts, but the nomadine bees are also remarkably diverse in size, making them an ideal model to test Harrison’s rule. Here, we reassess the phylogeny of Nomadinae, including the expanded sampling of the genus Nomada, to explore host shift fluctuations throughout their evolutionary history and test the applicability of Harrison’s rule for the subfamily. Our phylogenetic results are mostly congruent with previous investigations, but we infer the tribe Hexepeolini as a sister taxon to the tribe Nomadini. Additionally, the results reveal discrepancies with the traditional classifications of Nomada. Ancestral state reconstruction of host use indicates that, early in their evolution, parasites used closer relatives, before attacking less related groups later. Lastly, we confirm Harrison’s rule in Nomadinae, supporting that body size dynamics influence the host shifts of cleptoparasitic bees.